Naming the Groups: Developing a Stable and Efficient Nomenclature Author(s): John McNeill Reviewed work(s): Source: Taxon, Vol. 49, No. 4, Golden Jubilee Part 2 (Nov., 2000), pp. 705-720 Published by: International Association for (IAPT) Stable URL: http://www.jstor.org/stable/1223972 . Accessed: 02/06/2012 04:06

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Naming the groups: developing a stable and efficient nomenclature JohnMcNeill'

Summary McNeill, J.: Namingthe groups:developing a stable and efficient nomenclature.- Taxon 49: 705-720. - ISSN 0040-0262. The binomial system of botanical nomenclaturehas existed for almost 250 years, the principleof a taxon having a single correctname determinedon the basis of priorityof publicationwas formalizedalmost 150 yearsago, andthe type methodfor the applicationof scientific names of has had internationalacceptance for almost 75 years. In this historic time-frame,the achievementsof the past 50 years are outlined and the question posed as to whetherthe next 50 yearshold any prospectof change,and indeedwhether any changeis possibleor even desirable.The requirementof botanicalnomenclature to provide a stable, unambiguousreference system for plant information implies an inherent conservatismof rules and procedures--eventhe smallest change to the Code, however beneficialit maybe in general,is virtuallycertain to have some destabilizingeffect. Despite this truism,it is suggestedthat the next few years will see quite majorchange. One of the least of these may be the developmentof a separatespecialist nomenclature for communi- cation about major phyletic lineages, not so dissimilar in practice from the specialist nomenclaturecurrently in existence for the micro-variantsimportant in cultivatedplants. Bionomenclatureprovides the mechanism for communicationabout the elements of taxonomy-elements that generallyseek to reflect the greatestinformation on patternsof biodiversity.It will continue to communicatethe general informationcontent of taxa effectively, only if it evolves to take fuller advantageof the opportunitiesof the electronic age. The historicaltendency to improvethe rulesof nomenclatureby continuous"tinkering" with the Code needs to give way to a recognitionthat stability and simplicity are key requirementsof users of names (amongstwhom professionalbiologists are a relatively small minority),and that web access to authoritativelists will generallybe theirpreferred approachto answeringthe nomenclaturalquestions that arise in study and use of plants, animals and micro-organisms.To remainrelevant, the botanicalCode, like the bacterio- logical, and now, to a degree,the zoological,must provide mechanisms for the endorsement of nomenclaturallists thatrepresent the productsof soundscholarship. Key words:bionomenclature, communication, information, International Code of Botanical Nomenclature,nomenclature, PhyloCode, taxonomy.

Introduction It is particularlyappropriate to review progress and prospects in botanical nomenclatureas partof the celebrationof the 50th Anniversaryof the foundingof the InternationalAssociation for Plant Taxonomy (IAPT), because it is the foundationand activitiesof IAPT thathave done morethan anything else in the past 50 years to fashion the form and structureof botanicalnomenclature as it exists at the start of the 21st century. As outlined in the Editorialintroducing this series (Stuessy & al., 2000), IAPT was established at the VII International Botanical Congress in Stockholm in 1950, in large measure the brainchild of Lanjouw who played a leading role in rebuilding botanical nomenclature after the disruption of

'Royal OntarioMuseum, and Royal BotanicGarden Edinburgh. Mailing address: Royal BotanicGar- den, 20A InverleithRow, Edinburgh,EH3 5LR, Scotland,U.K. E-mail:[email protected].

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WorldWar II. The originsof IAPT have been well describedelsewhere (cf. Cowan & Stafleu, 1982; Stafleu, 1988), but it is worthemphasising some of the particularly nomenclaturalaspects of the contextin whichIAPT was born. Briquet, the first Rapporteur-gen6ral,and Rapporteur-gen6ralfor the Vienna (1905), Brussels (1910) and Cambridge(1930) Congresses (Briquet 1906, 1912, 1930), died in 1931 before the CambridgeRules could be completed(Harms, the Vice-rapporteur,and otherssuch as Sprague,who succeededBriquet as Rapporteur- g6neral,saw to its completionbut only just before the 1935 AmsterdamCongress). Althoughthere was a NomenclatureSection (then called a Sub-section)meeting and some importantdecisions were taken at Amsterdam,the vacuum left by Briquet's death and the gatheringstorm in Europe resulted in no Amsterdamcode being prepared.The so-called"Brittonia Rules" (Camp & al., 1947) publishedafter World WarII largelyfrom the notes of U.S. participantswere an unofficialrecord2, while a gathering hosted by Lanjouw in Utrecht in June 1948 of a group of nomenclaturalists,many of whom had been membersof the pre-war"Commission de Nomenclatureg6n6rale" that was the predecessor of the present General Committeefor Botanical Nomenclature,laid the groundworkfor the Stockholm meetingsand for IAPT (cf. Lanjouw,1950a, b). So IAPT is largelythe resultof Lanjouw'sdetermination that there would be an ongoing organizationto promotetaxonomy and nomenclatureinternationally and to ensure that the lacuna caused by Briquet's sudden death before completing the CambridgeRules would not occur again. Incidentally,one of Lanjouw's first actions,once it was clear thatthe Secretariatof the new IAPT would be in Utrecht, was to hire a relativelyrecent doctoralgraduate (his thesis on Vochysiaceaehad been defended in the week of the 1948 nomenclaturemeeting in Utrecht),just returnedfrom Indonesia, to do the day-to-daywork of runningthe Association.That person was, of course, Frans Stafleu. When we compare the efforts of Briquet (1930) to bringtogether the variousproposals to the CambridgeCongress, published -or not published-in a diversity of media, with the coherent publication, nowadays,of almost all nomenclaturalmatter in one of the two IAPT publications, Taxonor Regnumvegetabile, we can appreciatethe enormouscontribution made to botanicalnomenclature by Lanjouwand Stafleu-and theirsuccessors-through the establishmentand operationof IAPT. Fifty years may seem a long time, but in the historyof botanicalnomenclature it is not so very long, and the last 50 years are to be seen as a period of gradual evolution ratherthan one of decision on principlesor one of major innovation.I discuss this furtherbelow, but, in a sense, botanicalnomenclature today is not so very differentfrom what it was like 50 years ago-and this is not all due to the reactionarymood of the St. Louis Congress(cf. Greuter& Hawksworth,2000)! Why do I say this? Well, one must comparethe change over the past 50 years with that in the previous200 years. The binomialsystem was introducedin 1753, 247 years ago, the principle of a taxon having a single correct name determined on the basis of priority of publication was formalized around 150 years ago in the 1842

2Theonly official recordwas thatby Sprague(1950) which appearedin Lanjouw'sreport of the 1948 Conference (Lanjouw, 1950b) that was not published until around the time of the Stockholm Congress.

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StricklandCode for zoology (Strickland& al., 1843) and in Candolle's(1867) Lois for botany (English translation:Candolle, 1868). The type method for the applicationof scientificnames of plantshas had internationalacceptance for almost 75 years, being finally adoptedat the CambridgeCongress in 1930 (Briquet,1935) with the rapprochementbetween the AmericanCode (Arthur& al., 1907), which alreadyhad the type method,and the InternationalRules (Briquet,1912), which did not. This was fosteredby the IthacaCongress of 1926, at which no formaldecisions on nomenclaturewere takenbut which was an importantstep in the integrationof the two codes that was promotedmost effectively by the collaborationof T. A. Spragueand A. S. Hitchcock[cf. Duggar,1929 (Vol. 2: 1413-1597, "L.Taxonomy" & 1781-1782 "General Proceedings of Section L - Taxonomy"); Nicolson, 1991]. The significance of the type method as opposed to a circumscriptional concept and the historyof its adoptionin botanyare well describedby Moore (1998) and need not be repeated,although I will referto it furtherbelow. Unquestionably,these three developments,the binomialsystem, the principleof priorityand the acceptanceof the type method,are all much more fundamentalto bionomenclaturethan anythingthat has occurredin the past 50 years at least as regardsplant (and animal)names, thoughnot those of bacteriaas discussedbelow. That being said, I believe that there are a numberof fundamentalissues facing botanical nomenclature today that may stimulate major change. In order to understand these, I think we need to look at the purposes and principles to which I referred above.

Purpose of nomenclature Lanjouw (1950a), in his preparationfor the Nomenclature Section of the StockholmCongress where IAPT was born,included on the flyleaf a quotationfrom the distinguishedBritish mycologist Ramsbottom(1942): "Nomenclatureis the handmaiden of taxonomy, not the mistress"! But nomenclature serves much more than taxonomy. One of the opening statementsin Davis & Heywood's classic Principles of angiosperm taxonomy (1963) [p. 1.] is: "Biologists must know what organismsthey are workingwith before they can pass on informationabout them to other people-a function of taxonomywhich makes stability of nomenclaturean importantconsideration." Or to quote anotherstatement that is now almost a clich6 [whichin partmeans that I do not know who first said it]: "A plant'sname is the key to its literature".Any user of plantsneeds to referto plantnames, and any changes in plant names are disruptiveto an enormouscommunity outside of taxonomy, indeedoutside of biology. Bionomenclaturedid not startwith Linnaeus;the uniquephrase-name, the nomen specificumlegitimum, has a much longer history. Linnaeus'consistent use of the binomialby linkinga nomentriviale to the genericname, which we have set as the starting-pointof modem nomenclature,was simply a shorthandway of referringto species discriminated by their true name, the nomen specificum legitimum (Stearn, 1957: ch. 10). As such it was a major step in the process of separating nomenclature from taxonomy, making it clearer that the name is just a means of communication about distinguishable components of what we would now call biodiversity. In other words, nomenclature is a mechanism for unambiguous communication about the

Jubilee Series p. 179 708 TAXON49 - NOVEMBER2000 elementsof taxonomy.But whatare these elements?Since the 15t-16t centuriesin RenaissanceEurope, these have been the variouskinds of organismsthat were being recognizedand arrangedto some degreein an hierarchyof relationship.Aristotelian logic created a formal frameworkfor these that is expressed in the hierarchical arrangementwith which we are all familiar, and which has acquired the name "Linnaeantaxonomy", which is a bit of a misnomer,not that Linnaeus did not practice it, but that he certainlydid not introduceit. But although this logical formalizationmay be unique to the culture within which scientific taxonomy originated,a clear and sometimesquite well-developedhierarchy of "taxa"seems characteristicof all folk taxonomies,as Berlin, Breedlove and others (cf. Berlin 1972;Berlin & al., 1973;Berlin, 1974:ch. 2) have described. The Linnaeantrivial name and the type methodwere steps toward the modern dictumthat nomenclature is, or shouldbe, independentof taxonomy,but this is only true up to a point. Biological nomenclatureas expressedin the internationalcodes makesno assumptionsas to the methods,principles or purposesof taxonomy-save one, thatthe unitsof taxonomy,the taxa being named,are in some way nested,as in the hierarchyof ranksrecognized in the botanicalCode, thoughnot necessarilyso formalized. Whereas a cladogramcan always be expressed as a fully nested hierarchy,albeit with very many ranks, and the relationshipsamong its terminal elementsrepresented as an ultrametric,just as those of an hierarchicalclassification (cf. McNeill, 1980), the assumptionof a rankedhierarchy, even althoughthese ranks may be quite arbitrary(cf. Stevens, 1997), may be inconvenient for naming individual clades, particularlyif these are being considered individually or are overlapping. Hence, although the internationalrules of nomenclatureare as appropriatefor namingcladistically defined groups as any others, they may prove inconvenient,or some might say inappropriate,for the namingof individualclades per se, and it is this that has led to the PhyloCode(Cantino & de Queiroz,2000). WhenI first met and listenedto Kevinde Queirozat a symposiumorganized by Jim Reveal at Marylandin 1996 (de Queiroz, 1996; Reveal, 1996), I suggestedthat his issue was a taxonomicnot a nomenclaturalone; he thoughtnot, but, in fact, we are both right. As the existence of the draft PhyloCode shows, there is indeed a nomenclaturalcomponent, but it is one thatstems directlyfrom a questionabout the principlesand purposes of taxonomy. If taxonomy,as the derivationof the word implies, is aboutclassification, about grouping organismsinto successively more inclusive classes, on whateverbasis, then the well-establishedinternational systems of nomenclature,however imperfect they may be, are applicableand appropriatemedia for naming. Or, put in another and, I believe, betterway: if the elementsof taxonomyare those that seek to reflect the greatest informationon patternsof biodiversity,then the existing systems of nomenclaturewill permit ready summarizationof this informationin a nested framework,communicating the general informationcontent of taxa effectively. If, on the other hand, names are sought for communication about lineages, without a nested hierarchy, a totally new system may well need to be devised, and the PhyloCode is evidently a first essay at such a system. This being said, I, personally, wonder whether communication about clades needs such an elaborate mechanism; would PhyloCode rules make the usage of "" or "eumagnoliids" in, for

Jubilee Series p. 180 TAXON49 - NOVEMBER2000 709 example, Soltis & al. (2000) any easier to understand?Although I realise that the proposersof the PhyloCodehave granderambitions, the namingof groups outside of the generalsystem of biology is not uniqueto clades. Most biologists recognize that much infraspecificvariability is not satisfactorilyaccommodated in detailed Aristotelian hierarchies,such as Ascherson & Graebner(1896-1938) for wild populationsor Mansfeld(1959) for such cultigensas Brassica spp. or Beta vulgaris. And in the case of cultigens, where it is importantto name fine details of infra- specific variation for commercial purposes, a separate set of rules has been developed, the International Code of Nomenclature for Cultivated Plants (Trehane & al., 1995). Its limitedhierarchy of ranks(cultivar and cultivar-group)are linkedto the hierarchyprescribed in the ICBNat whateverlevel is appropriate,sometimes to species, but more commonly to . It may be that botanistsshould follow the exampleof zoologists and not seek to regulatethe namingof groupsabove the rank of family. Those who wish to use groupsdefined by a single type as providedunder the ICBNcould continueto do so, but othersmight want to experimentwith groups defined on inclusion or exclusion of several elements or on particularcharacter distributionas proposedin the PhyloCode. It is not, however, the purpose of this paper to evaluate the PhyloCode (such commentariesare proliferating,cf. Moore, 1998; Redhead,2000; Withgott,2000; Benton,in press),or even to compareit with the existingcodes. I did, however,want to put it into its taxonomicand nomenclaturalcontext, and I will come back to that in the conclusions.

Principles of nomenclature I have already referred to the principle that nomenclature should be as independentof taxonomyas possible and how this has been progressivelyachieved by, for example, the introductionof the binomial (replacingthe diagnostic poly- nomial), and the adoptionof the type method for determiningthe applicationof names.I do believe thatthis has been a beneficialtrend in facilitatingstability in the applicationof names without significantdiminution in clarity of communication. This being said, it is not the only routethat could be taken,and it is not always the one thatis most beneficialfor communication.For examplethe names Leguminosae and Papilionaceaeare immediatelymore informativethan is the name Fabaceae, and althoughthese namesand theirusage are enshrinedin Art. 18.5 of the botanical Code (Greuter& al., 2000), theirrelative application is defined circumscriptionally ratherthan by the type, which is the same for all three.In a similarvein, Berendsohn (1995) in developing the concept of "potentialtaxa" recognized that information storagein databasesto be unambiguousneeds to be linkedto the usage of a nameby a particularauthor and not simply to the type of that name--essentially a returnto the circumscriptionalcitation procedurethat was common and informativein the 19th century,but that would seem an intolerableburden in literaturecitation today, but perhaps not at all a burden in database management. But the key point that I would make on the principles of nomenclature is that its function is to serve taxonomy. This has two aspects to it: if the taxonomy of a particular group is in a state of flux, the nomenclature will be also; nomenclatural rules cannot solve the problems of taxonomy. But it also means that there is no right

Jubilee Series p. 181 710 TAXON49 - NOVEMBER2000 or wrong in the formulationof nomenclaturalrules, although, of course, there generallyis in their application.The only issues are simplicity and clarity and the provision of stability. Nothing in nomenclatureneed be sacrosanct. Even the principleof priorityof publication,adopted throughout bionomenclature since the middle of the last century,is simply a convenienttool to determinein an unbiased manner which of two or more competing names should take precedence for a particulartaxon. When applying the principle is disruptive to nomenclatural stability,it should be readilyabandoned, a position well demonstratedby the very many proposalsthat have been accepted over the years to conserve widely used namesover earlierlittle used ones, or else to rejectthe latterentirely (nomina utique rejicienda).Priority, of course, remainsthe appropriatetool to determinecorrect usage when two taxaare mergedand different botanists, sometimes in differentparts of the world,would be discomfitedwhichever name is adopted,as appearsto be the case with the merger of the African caesalpiniaceousgenus ColophospermumJ. Lonard with the IndianHardwickia Roxb. (cf. Brummitt,2000). Likewise,if a type turns out to be taxonomicallydifferent from what taxonomists have generally presumedit to be, it shouldbe changed(in the mannerprovided by Art. 14.9 of the Code; Greuter& al., 2000), and nomenclaturalconfusion thus avoided by main- taining existing usage of the name, as is often done, e.g., for that of Filicites heterophyllusBunge [the ultimatetype of Neuropteris(Brongn.) Sternb.] which was found to be referableto a differentspecies from thatto which the name is currently applied (Skog, 1999). On the other hand, maintenanceof established types is appropriatewhen thereare significantlosers as well as winners,as in that proposed as a resultof the restrictionof Myricato its type, the EurasianM. gale L. (and one otherspecies), requiringa genericname change for over 40 Africanspecies hitherto includedin Myrica(cf. Brummitt,1999). This emphasison practicalityrather than inviolate principleis supportedby the recognitionthat in other nomenclaturalsituations the various Codes, notably the botanicaland zoological, historicallyhave adopteddifferent ways of resolving the same question. Some of these may be simpler to understandand communicate, which may make them preferable,others may be logically more satisfyingin terms of otherprovisions of the Codes, whereassome may have an instabilitycomponent thatmight argue against them, but they are all operational.For example,of the three logical ways to determinethe correctname when a species is transferredfrom one genus to another (cf. McNeill, 1997), probablythe simplest to understandand communicateis the one that no currentCode adopts, namely that of the so-called "Kew Rule", in which the earliest name, i.e. binomial,adopted in the second (or indeedany) genus is correct,i.e., becomesthe acceptedname. On the otherhand, the requirementof the zoological Code (Ride & al., 1999) that the earlier published epithetbe used regardlessof its priorindependent use withinthe second genus (i.e. thatto which transferis being made)creates the possibilityof instabilityin the name of a taxon, the one already bearing that epithet in the second genus, that is not involved in the taxonomic change-the situation of "secondary homonymy". For example, if Lychnis sibirica L. (Sp. Pl. 436. 1753) is transferred to Silene, then under the zoological Code, Silene sibirica (L.) Pers. (Syn. Pl. 1: 497. 1803), based on Cucubalus sibiricus L. (Syst. Nat. ed. 10. 2: 1031. 1769), would require a new

Jubilee Series p. 182 TAXON49 - NOVEMBER2000 711 name, whereas, under the botanical Code, L. sibirica is given the next earliest available epithet (and probably becomes S. samojedorum), and the name S. sibirica is retained for the species that is not involved in the taxonomic change. Conversely, the zoological principle of "co-ordinate status" in the rank groups (family, genus and species) seems to facilitate the preservation of names or epithets within those groups in a way that the botanical principle of complete rank-limited priority does not. But there is no right or wrong. Nomenclature is there to serve the user community and it is important that the rules are used to facilitate this-and, as the history of the past 50 years has demonstrated, that they be modified if they cannot facilitate adequately.

The past 50 years In his introduction to the Stockholm Code, Lanjouw (1952) expressed the "hope that we will not change our Code at every subsequent Congress." How false a hope! There have been eight Codes published in the 48 years since then, one for each of the eight Congresses. At some Congresses the rules changed very little; at others there was more substantial change or at least substantial change in the form and structureof the Code, e.g., in the numbering of paragraphs,in the merging of certain appendices into the main body of the Code, or dropping them altogether. Many of the changes to the Articles of the Code were very important for particular situations, for example those dealing with Art. 59 on fungi with pleomorphic life cycles at Sydney, and those on fossil morphotaxa (form-taxa, organ-taxa) at St. Louis, but none was comparable in significance with the introduction of the type method at the Cambridge Congress of 1930. A detailed history of the changes to the rules over the past 50 years has yet to be written-Nicolson's (1991) excellent history of botanical nomenclature essentially ends with the Paris Congress of 1954 -and this is not the place for such a treatise. In passing I would comment, however, that the budding historian of plant nomenclature should be grateful for the innovation for post-war Codes made when Ed Voss was Rapporteur-g6n6ralin that the Preface to the Sydney Code (Voss & al., 1983) outlines the main changes in that edition from previous ones, a tradition that has been maintained ever since, but something that neither Lanjouw nor Stafleu attempted, although comparisons had been extensive in pre-war Codes, and summaries or more detailed accounts had sometimes appeared elsewhere (e.g., Lanjouw, 1951; Stafleu, 1964b). Over these 50 years there has been, however, an almost continuous debate that reached what was probably its final fruition at the Tokyo Congress of 1993. This was the issue of conservation and/or rejection of species names. The principle of conservation of names goes back to the very beginning of the international rules, because in response to the work of Otto Kuntze in his Revisio genera plantarum (Kuntze, 1891-98) who, in applying strict priority, threatened the change of over 500 familiar generic names, the Vienna Congress approved a list of nomina generica conservanda (Briquet, 1906, cf. also Nicolson, 2000). (This was not, however, a feature of the American Code, as it was anathema to N. L. Britton, one of its founders, which is perhaps why the concept has never had the support in North America that it has in Europe; cf. McNeill & Greuter, 1987). At the Stockholm Congress in 1950 there was a proposal to introduce nomina specifica conservanda (though not the first such, which dates to 1929, cf. Nicolson, 2000). This proposal

Jubilee Series p. 183 712 TAXON49- NOVEMBER2000 was rejected despite pleas by John Gilmour, then Director of the Royal Horticultural Society's Garden at Wisley, that not to accept it would be extremely troublesome for horticulturists and other general users of plant names. The alternative strategy of nomina specifica rejicienda, thought to have greater chance of success as involving a smaller case-load and still dealing with some of the most serious problems, was canvassed, e.g., by George Taylor, as early as 1954 at a conference (Stafleu & Lanjouw, 1954) similar to the 1948 one, involving a select group of nomen- claturalists preparatoryto the Paris Congress. These initiatives had their supporters, and other initiatives to minimize name changes were undertaken(cf. Stafleu, 1964a), but no breakthroughcame until the Sydney Congress in 1981. It was one particular dramatic case that persuaded even that arch-opponent of any departure from strict application of the rules, Art Cronquist (New York), to lend his and with it much North American support for a very limited proposal to permit the conservation of names of species "of major economic importance". The cause clelbre was, of course, that of the scientific name for common bread (or soft) wheat. For several decades this had been understood to be Triticum aestivum L. and, indeed, this name exemplified the article of the Code dealing with choice between names of equal priority, Fiori & Paoletti in 1896 being thought to be the first to combine T. aestivum L. (1753) with T. hybernum L. (1753), choosing the former. The discovery by Kergu6len (1980) that, in 1821, M6rat had done the same thing but chosen Triticum hybernum would have meant a change, for purely nomenclatural reasons, in the scientific name of a major economic crop, something that few could justify. And so there was the limited adoption of nomina specifica conservanda at the Sydney Congress in 1981 (cf. also Hanelt & al., 1983). [As a personal aside, I may say that the form of words ("major economic importance") that would secure the votes Art Cronquist carried were those we worked out together while I was his guest at a delightful dinner in a sea-food restaurant overlooking Sydney Harbour, a view familiar to those who watched the recent Olympic Games. But not even this restrictive clause was sufficient to convince others, such as Ray Fosberg (US), of the wisdom of what seemed to them such a radical step. Ray was most vexed that, being then newly retired, he was no longer carrying the Smithsonian votes, particularly as his successor, Dan Nicolson, was fully persuaded, and also as the proposal gained its 60% majority with a plurality of only four!] Formal procedures for the rejection of names that had been widely used in a sense not including their type were also established in Sydney, while the provisions for nomina specifica conservanda were slightly extended at the Berlin Congress, but it was not until Tokyo that the present provisions for conservation of the name of any species, genus or family "in order to avoid disadvantageous nomenclatural changes" were enacted. At the same time it was made possible to propose for rejection any name (i.e., at any rank) that would cause such disadvantageous change. While the work-load arising from this widening of provisions for conservation and rejection has been very considerable, I have no doubt that this has been most worthwhile in permitting nomenclature to serve the needs of users more effectively. One need only look at the voting patterns of the Permanent Nomenclatural Committees for particular groups to see that, although there are a small number of extremely troublesome cases upon which the Committee involved is very divided,

Jubilee Series p. 184 TAXON49 - NOVEMBER2000 713 the vast majority of proposals are either supported almost unanimously or rejected almost unanimously. In other words most proposals are seen to be good and desirable, a number are clearly inappropriate, and there is the inevitable residue of problem cases. There can be no doubt that the decisions of the Tokyo Congress make it possible under the rules not to displace "well established names for purely nomenclatural reasons, whether by change in their application or by resurrection of long-forgotten names" to quote from a resolution of the Congress (cf. Greuter & McNeill, 1994). Although not even the General Committee for Botanical Nomenclature has the supreme plenary powers over the application of the botanical Code that the International Commission for Zoological Nomenclature has over those of the zoological Code, there is no doubt but that the ICBN does now provide the tools to minimize nomenclatural change, always provided that individual botanists and the Permanent Nomenclature Committees take time and make the effort to use these tools and interpretthem wisely. What of the other achievements and nomenclatural initiatives of the past 50 years? I have already referred above to some improvements and clarifications in particularissues, and this has been the general trend. I think it is fair to say that the Code is now a much more precise document than it was after Stockholm, giving clearer instruction in many problematic areas, though whether this makes the document an easier one to use is an open question. I may be biased but I believe that in at least one area the Code remains hopelessly confused. Despite the extensive clarifications on orthography approved at the Leningrad Congress in 1975, this portion of the Code remains problematic-it attracted some 70 proposals for amendment in St. Louis, almost one third of the total (215). I would venture to suggest that the problem of orthography is an intractable one, at least in so far as dealing with it through amendments to the Code, and this is alluded to below. Although orthography seems to me the most tricky, there are other areas of ambiguity in the Code, such as criteria for effective publication (and its relation to electronic publication) that cannot readily be resolved by rewording the Code. But what of the other three initiatives that provoked so much controversy and negative comment at St. Louis: the Draft BioCode, Names in Current Use (NCUs), and Registration? The first can be dealt with simply. As was always understood by those involved in its development, the Draft BioCode (Greuter & al., 1998) was never intended to replace existing Codes in contemporary nomenclature; it was always a code for the future (cf. Greuter, 1996, in press; McNeill, 1996) and, despite the concerns expressed by many, both before (Orchard & al., 1996) and during the St. Louis Congress, there were no proposals on the BioCode, per se, presented to the Nomenclature Section in St. Louis, nor would any have made sense unless other initiatives had been very much further advanced. Some of the communication issues between codes of nomenclature that development of the Draft BioCode made apparent are important and continue to be important-to understand, for example, that botanists and zoologists mean something totally different by the words "valid" or "available". Harmonization of terminology that will clarify understanding and not destabilize names remains a desideratum, whereas any significant changes to the Codes themselves have always been recognized as impossibly destabilizing. The

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other two issues, approved lists and registration, have much wider implications and are alluded to below in the context of botanical nomenclature in the future.

The present and the future I am sure that many of us grew up with the dictum that, while applying a set of rules governing the names of plants might lead to some initial changes in familiar names, as the Code was applied universally these would become fewer and fewer to the point of vanishing. As the history of the past 50 years demonstrates, and as I think most taxonomists now recognize, this ideal situation does not work out quite that way in practice-and not only because of changing taxonomic concepts. More thorough typification in accordance with the rules is one major cause of potential change in the application of names, e.g., Redhead & al. (in press) have demonstrated that, under strict application of the rules, the type of Psathyrella (Fr.) Qu6l., a name currently applied to a genus of well over 100 species, is referable to Coprinus sensu stricto. Another is the continuing discovery of early nomenclatural acts that have been overlooked, the most dramatic case being probably that described above, namely the discovery of the adoption by Merat in 1821 of Triticum hybernum L. (1753) rather than T. aestivum L. (1753) for common wheat (Kerguelen, 1980). A third is represented by the unwanted by-products of otherwise apparently desirable changes in the Code. For example, the ambiguity created by Necker's "unitary designations" resulted in these being disallowed as generic names by the Code back in 1961. It was realised only a few years ago that, as a result, the familiar composite generic name lost its priority and a conservation proposal has been made (Robinson & Funk, 2000). Finally, it is still the case that many hitherto overlooked publications are being found to contain nomenclaturalnovelties that have never been indexed; this is largely in the so-called "grey literature" of early horticultural journals and catalogues and the like, often with minimal validation of names, but often nomenclaturally destabilizing. One such example is that of Arundinaria nitida "accidentally" published in an editorial report of a meeting (cf. Stapleton & Zijlstra, in press). This realization that we will never be free of name changes "for purely nomenclatural reasons" has had two beneficial effects: more extensive provision for overriding the rules, and less enthusiasm for "tinkering with the Code". Even although the St. Louis Congress declined to incorporate stability of names as an overriding principle in the Code, even those opposed to the new principle made clear that they whole-heartedly supported stability of names, something already manifest from the Tokyo Congress by the adoption of procedures to reject any name that would cause "disadvantageous nomenclatural change" and, likewise, to conserve the name of any species, genus or family for the same reason. This has not always been the case. Far too often, when troublesome nomenclatural implications arose from hitherto overlooked or rejected publications or from a strict application of the rules, the temptation to amend the Code to deal with the specific issue has not been resisted. Sometimes, of course, this was because there was no alternative mechanism under the Code (e.g., before conservation of species names was permitted), but not always. For example when Reed Rollins and other members of the then Committee for Spermatophyta discovered that accepting the first lectotypifications by Britton

Jubilee Series p. 186 TAXON49 - NOVEMBER2000 715 and Brown would lead to very confusing change in the application of many familiar Linnaean generic names, particularly in Brassicaceae, an ill-phrased proposal was accepted that would allow American Code lectotypifications to be superseded on the questionable ground that they were made "arbitrarily" (when very many lectotypifications made under the international Code are even more arbitrary)or, as it later became in the Sydney Code (Voss & al., 1983), by a "largely mechanical method of selection". Better either to have moved the starting date for priority of lectotypification to a date at or after the introduction of the type method into the International Rules, or else deal with each case individually by conservation. Fortunately it is now almost universally realised that any change in the provisions of the Code, no matter how small, is certain to have some nomenclatural impli- cations that will not necessarily have been thought through by the proposers or even detailed in the Rapporteurs' comments. Whereas the provisions for conservation and rejection can mitigate some of these negative effects, it rarely makes sense to try to amend the Code to deal with highly specific situations. This does not mean that the Code cannot be modified or improved in certain areas, but it needs to be done with great care. Certainly "tinkering" with the Code is not generally a prescription for stability of names. There are two realities that I believe will have a profound effect on nomenclature in the future, whether we like it or not. The first is that electronic access to information, particularlyfactual information such as that relating to scientific names, has effectively become the norm--even I, working in a major taxonomic institution with an excellent library, find it far simpler to look at the IPNI website (http://www.ipni.org) from the computer on my desk than go down to the library to check the IK CD-ROM, far less the bound copies with their 20 supplements. (Cf. also the discussion of current name-based web applications by Pennisi, 2000, and Bisby, 2000.) Moreover when the consumer cannot get such information reliably from a website, he or she becomes somewhat irritated and may go for the best that can be obtained electronically, regardless of its quality or its conformity with international rules of nomenclature. Linked to this is the reality that the number of persons who have both time for nomenclatural work and access to traditional taxonomic literature is steadily declining. Moreover, this population is being increasingly taxed by the very welcome, but still labour-intensive, process of conserving and rejecting individual names. Just as it is generally more cost-effective to tackle problems with particularnames directly, than attempting to do so through amendments to the Code, for which there is likely to be payment "down the road", it is even more the case that endorsing lists of names developed through sound scholarship and making these widely available electronically, is likely to be the only practical solution for nomenclatural stability in the future. This is essentially what the bacteriologists did in the 1978 edition of their Code, in which they established a list of Approved Names, which effectively set 1980 as a new starting date for names of bacteria. This sought to take account of all bacterial names in taxonomic use at that time. It in no way restricts the application of new names as a result of taxonomic research, but it does liberate bacteriologists from worrying about the application of other names from the past and allows them to consider only that restricted number whose application is established and

Jubilee Series p. 187 716 TAXON49 - NOVEMBER2000 documented. It is encouraging that zoologists have incorporated provisions to endorse lists of names in their recently published revised Code (Ride & al., 1999). In botany, the Names in Current Use proposals, which had some resemblance to the bacteriological initiative, were, as noted above, very unpopular in some circles. I suspect that this was more because of the quality of some initial draft lists than because of the fundamental principle, which, after all, has been in the Code since the Montreal Congress in the form of the Appendix of names of families of Spermatophyta (App. IIB in recent Codes). Some opposition may have stemmed from the misconception that such lists would constrain taxonomy, which is totally unfounded. What does seem to be the case, however, is that the timing of the NCU proposals was ill-advised. It is instructive to consider how two of the major changes in the principles of nomenclature came to be adopted in botany. The first, of course, was the Linnaean binomial system; neither Linnaeus nor anyone else established a rule that names should be in the form of binomials. Instead the first, and great synthesis of the world's flora, Species plantarum (Linnaeus, 1753), quickly recognized as such a synthesis, happened to include in the margins these single word epithets that could so conveniently be linked to the generic name. A vitally important work that everyone had to use incorporated the system and it became adopted almost immediately (barring a few hold-outs). Much the same happened with the introduction of the type method into the International Rules. I referred above to the collaboration between Hitchcock and Sprague at and following the Ithaca Congress of 1926. This was developed in very practical ways and extended to involve Sprague's assistant, and later wife, M. L. Green, in the production of lists of "standard species". This was a diplomatically well-chosen phrase, as "type", a feature of the schismatic American Code that would be anathema to many Europeans (just as conservation of names, then peculiar to the International Rules, was to many Americans). I am sure that there was no way that the type concept would have been accepted by Europeans into the International Rules but for the work of Hitchcock and Green in demonstrating what its implications might be. By publishing "standard-species" of all Linnaean generic names (Hitchcock & Green, 1928) and of all conserved names (Green, 1928), even if, in the event, these were never formally accepted either at Cambridge or Amsterdam, I am certain that they reassured many European botanists. [As another aside, although much too young to know of these events at first hand, I did attend the Edinburgh Congress in 1964 and, seeking a cup of coffee in Edinburgh's gracious Georgian Assembly Rooms that served as a Congress club, chanced to join a very spry old lady who turned out to be none other than Mrs. Sprague (M. L. Green), who, I think, was ratherpleased that someone as junior as I had heard of her contributions to botanical nomenclature. Her achievements were especially noted at the Nomenclature Section dinner of that Congress-apparently the first time this now traditional event was held (Stafleu, 1964b)]. I believe the experience of the acceptance of binomials and of the incorporation of the type method is suggestive of the route forward. What we need are more and more authoritative lists, readily available to users, which, as they become more complete and hence more valuable will, like Linnaeus's binomials, become de facto and then inevitably de jure standards for plant nomenclature. They will also provide

Jubilee Series p. 188 TAXON49 - NOVEMBER2000 717 benefits that amendmentsto the Code seem incapable of achieving, such as standardizationof orthography,and proof of valid publication,and will be the catalystfor authorsto ensurethat their new names are listed-"registered",if you like. However,whatever nomenclaturally inclined taxonomists decide in our sexen- nial meetings,lists will be producedto meet user needs. It will be much betterfor unambiguouscommunication and long-termstability of names if these lists re- presentsound scholarship and application of the rulesof nomenclature.

Conclusions In summary,I see a future that as evolutionarybiologists we should readily understandand appreciate.Changes in technology and changes in methods of working will lead to changes in our nomenclaturalneeds. It will be those nomenclaturalcomponents that meet these needs that will survive and develop. Those thatdo not will atrophy,much as the nomenspecificum legitimum did before the onslaughtof the binomial,or the rightto select a new namefor a new taxonomic conceptbefore the entrenchmentof the type method. It is always difficultto predictthe paththat evolutionwill take and the futureof nomenclatureis no exception. It is possible that a need will be found for communicationabout major evolutionarylineages for which a non-hierarchical system of nomenclature,such as that suggestedin the PhyloCode,is indicated.On the otherhand, I do not see any evidence foreshadowinga decline in the need for a general reference system for biodiversity, not only at the specific level and significantcomponents of variationbelow that, but also with a substantialhierar- chical componentabove it, covering,in other words,those aspects of nomenclature with whichthe currentinternational codes are most involved. But this will be in a scientificmilieu in which internetaccess for informationhas become much more pervasivethan it currentlyis, and where contemporaryauthor- itative lists ratherthan referenceto extensive older literaturewill be seen by most scientistsas the solutionto nomenclaturalquestions. To remainrelevant to science, it will have been imperativefor the internationalcodes to have respondedto this need by lendingtheir weight and authorityto lists that representsound scholarship, paving the way for, and eventually achieving, new starting dates for plant bionomenclaturecomparable to thatachieved by the bacteriologistsin 1980.

Acknowledgements I am greatlyindebted to F. R. Barrie,W. Greuter,G. Moore,D. H. Nicolson,P. F. Stevens, P. Trehane,and J. Wiersemafor commentsand suggestionson earlierversions of this paper, and for correctionsto them. The opinions expressed remain, of course, the author's.

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